Bornean Keeled Pit-viper

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Life from Headwaters to the Coast

Biodiversity Between Land and the Sea


Edited by

Jayasilan Mohd-Azlan Mohamad Kadim Suaidi Indraneil Das

Edited by Mohd-Azlan, Suaidi & Das



Biodiversity Between Land and the Sea


Life from Headwaters to the Coast


Edited by

Jayasilan Mohd-Azlan Mohamad Kadim Suaidi

and Indraneil Das

Biodiversity Between Land and the Sea



Bako: Biodiversity Between Land and the Sea


Published by UNIMAS Publisher Universiti Malaysia Sarawak 94300 Kota Samarahan Sarawak, Malaysia.

Website: in association with

Natural History Publications (Borneo) Sdn. Bhd. (216807-X) A913, 9th Floor, Wisma Merdeka Phase 1

P.O. Box 15566, 88864 Kota Kinabalu, Sabah, Malaysia.

Tel: 088-233098 Fax: 088-534502 Website: Life from Headwaters to the Coast:

Bako. Biodiversity Between Land and the Sea

Edited by Jayasilan Mohd-Azlan, Mohamad Kadim Suaidi and Indraneil Das ISBN 978-967-0054-16-2

First published 2023.

Copyright © 2023 UNIMAS Publisher and Natural History Publications (Borneo) Sdn. Bhd.

Photographs copyright © 2023 with respective photographers.

Copy Editor: Genevieve V. A. Gee

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of the copyright owner.

Half-page: Sea Stack. Photo: Hans Hazebroek Front cover: Silvered Langur. Photo: Chien C. Lee

Frontispiece: Sandstone gate at Telok Tajor. Photo: Hans Hazebroek Printed in Taiwan

Perpustakaan Negara Malaysia Cataloguing-in-Publication Data

Life from Headwaters to the Coast : BAKO : Biodiversity Between Land and the Sea / Edited by Jayasilan Mohd-Azlan, Mohamad Kadim Suaidi and Indraneil Das. ISBN 978-967-0054-16-2 1. Biodiversity--Malaysia--Sarawak.

2. National parks and reserves --Malaysia--Sarawak.

3. Taman Negara Bako (Sarawak, Malaysia).

4. Government publications--Malaysia.

I. Jayasilan Mohd-Azlan. II. Mohamad Kadim Suaidi. III. Indraneil Das.



Bako: Biodiversity Between Land and the Sea Foreword ... vii

by YB Dato Sri Haji Abdul Karim Rahman Hamzah

Preface ... ix Introduction ... 1 Introducti

General Section

Reminiscences of Bako ... 3 by Gathorne Cranbrook

Geology and Geomorphology ... 5 by Hans Hazebroek

Floral Accounts

The Vegetation ... 19 by Meekiong Kalu, Stephen Teo Ping, Mohd Effendi Wasli, Nursafinas Jelani, Mohd Akmal Mohd Raffi and Yazid Kalbi Two Endemic Herbs ... 25

by Wong Sin Yeng

Carnivorous Plants ... 29 by Chien C. Lee

Mushrooms ... 35 by Mohamad Hasnul Bolhassan, Mohd Haqim Mohd Haizar Nazry, Nur Atiqah Asiqin Sulaiman Khan, Mohamad Nurfazillah Mohamad Ramzie Faizal and Fatimah Daud

Faunal Accounts

Mosquitoes ... 37 by Wong Siew Fui, Ichiro Miyagi, Takako Toma and Takao Okazawa Dragonflies and Damselflies ... 41

by Rory Dow and Graham T. Reels Butterflies ... 47

by Pang Sing Tyan, Ratnawati Razali and Wan Nurainie Wan Ismail

Land Snail ... 53 by Mohd Zacaery Khalik and Mohammad Effendi Marzuki Crabs and Shrimps ... 57

by Jongkar Grinang

Inland Fishes ... 60 by Kelvin K. P. Lim

Amphibians ... 65 by Indraneil Das, Ramlah

Zainudin, Pui Yong Min, Elvy Quatrin Deka and Taha Wahab Reptiles ... 69

by Indraneil Das, Pui Yong Min, Sabariman Hassan and Taha Wahab Birds ... 77

by Mohamad Fizl Sidq Ramji, Nurul Ashikeen Ab Razak, Ivana Berlinda Bilang, Rahah Mohd.

Yakup, Isa Sait and Badiozaman Sulaiman




Bako: Biodiversity Between Land and the Sea


Bats ... 91 by Faisal Ali Anwarali Khan, Muhd Amsyari Morni, Julius William-Dee, Norfarhana Mazlan, Wan Nur Syafinaz Wan Azman, Emy Ritta Jinggong, Nur Afiqah Aqilah Azhar, Roberta Chaya Tawie Tingga, Mohd Ridwan Abd Rahman and Isham Azhar

Non-volant Small Mammals ... 95 by Faisal Ali Anwarali Khan, Julius William-Dee, Muhd Amsyari Morni, Nor Al-Shuhadah Sabarudin, Muhammad Amin Iman Azmi, Syamzuraini Zolkapley, Roberta Chaya Tawie Tingga, Mohd Ridwan Abd Rahman and Isham Azhar

Large Mammals ... 99 by Mohd-Azlan Jayasilan and Sally Soo Kaicheen

Species Highlights

Saltwater Crocodile ... 106 by Engkamat Lading and Indraneil Das

Bornean Green Pit-viper ...111 by Veronica Martin and Indraneil Das

Proboscis Monkey ... 114 by Bolhan Budeng, Joan

Bentrupperbäumer and David Gillieson

Silvered Langur ... 117 by Bolhan Budeng, Joan

Bentrupperbäumer and David Gillieson

Sunda Colugo ... 120 by Dzulhelmi Nasir and

Nursyereen Mohd Nasir

Bearded Pig ... 123 by Mohd-Azlan Jayasilan and Sally Soo Kaicheen

Human Dimensions

Ecotourism ... 125 by Arianti Atong, Abang Norizan Abang Median, Jalani Kelat and Suhaili Mokhtar

Contributor Affiliations ... 131


Bako: Biodiversity Between Land and the Sea



arawak retains some of the richest biodiversity in the world. It is home to many endemics and species of conservation importance. Some of the best examples can be found in the State’s extensive network of protected areas. Many of us here in the Ministry continuously explore the exquisiteness of biodiversity in the hopes of harnessing and sharing of information with the general public, to appreciate such elements present in our protected areas.

This book represents but a sample of the work done by academics in the realm of biodiversity from Universiti Malaysia Sarawak and experts from various other agencies. I would like to commend

the efforts by these researchers who supported us in collecting information on the biodiversity in such species-rich areas as Bako, which forms the material for the book.

The work is also expected to be important for local communities, to enhance their understanding, appreciation and perhaps eventually, guide their use of such resources sustainably, acting as an interpretation tool to guide ecotourists and naturalists.

As will be evident to the readership, a variety of approaches have been taken by the authors of the volume. Sections, starting with reminiscences from the early days by the Earl of Cranbrook, and on geology and geomorphology, are divided along taxonomic and thematic lines. These include a general account of the tree flora and selected herbaceous flora, a review of carnivorous plants and one on the mushrooms. The faunal accounts include both invertebrates and vertebrates, ranging from mosquitoes to monkeys. A section highlights the biology of Bako’s charismatic species, that attract so many tourists to the Park. Finally, the section on human dimensions round up the volume, with a chapter on ecotourism in Bako National Park.



Bako: Biodiversity Between Land and the Sea

It is my hope that this book will contribute in a signifi cant way by encouraging more people to appreciate nature, explore our biodiversity and win more supporters. I anticipate that this volume will be useful to stakeholders to whom we remain connected through our common views on biodiversity conservation for the future generation.

Yang Berhormat Dato Sri Haji Abdul Karim Rahman Hamzah Minister of Tourism, Creative Industry, Performing Arts;

Minister of Youth, Sports & Entrepreneur Development Sarawak FOREWORD


Bako: Biodiversity Between Land and the Sea


Veronica Martin and Indraneil Das


he Bornean Keeled Pit-viper (Tropidolaemus subannulatus) is a strictly arboreal, venomous snake that is commonly encountered in Bako National Park. It possesses a heat-sensitive pit behind the nostrils, the source of its English common name. The species is widespread across many of the major islands of south-east Asia, including Borneo (Kalimantan, Indonesia; Labuan, Sarawak and Sabah, Malaysia and Brunei Darussalam), and the southern parts of the Philippines. Besides Bako, the species can be seen along the sandstone hills of Kubah National Park and Gunung Santubong National Park, Lambir Hills National Park, Ranchan Pool forests of Serian, Gunung Penrissen, the alluvial forests of Gunung Mulu National Park, the limestone hills of Bau, the peat swamps of Kota Samarahan, and many other localities across Borneo.

Characteristics of the species include a fl at-topped and steep-sided head, in addition to its distinctive triangular shape, the rear of head wider than the neck, the forehead scales keeled, the tail prehensile or capable of gripping small branches and other objects and extreme dimorphism or differences

Fig. 1. Adult female (left) and male (right) Bornean Keeled Pit-viper. Bako: Biodiversity Between Land and the Sea



Bako: Biodiversity Between Land and the Sea


between adult male and female.

Similar to nearly all pit-vipers, this species is ovoviviparous, whereby eggs hatch within the mother’s body. As juveniles, males and females are similar in morphology and colouration, both having a slender body, and therefore hard to distinguish. With growth, males and females become sexually dimorphic as well as dichromatic:

males are relatively more slender, whereas adult females become heavy-bodied, longer and heavier, reaching up to 1 meter in length.

Females also show greater variation in colouration: some are yellow-green, with bluish crossbands, while other turn blue- black, with small yellowish green dots arranged in a pattern forming crossbands. This contrasts with adult males which show a pale green ground colour, with rows of bright red spots on each side of the flanks; these spots either fade with growth or turn blue. The bright green and red colouration helps them blend with their leafy habitat,

critical for an ambush predator, and also presumably in escaping predators.

Another interesting adaptation shown by many green pit-vipers is a bright tail-tip, thought to be used as a lure to attract prey.

Being associated with trees and other forest vegetation, the Bornean Keeled Pit-viper is a species of lowland dipterocarp forests. Movements are more typically vertical, some ascending tree canopies and adults are known to move between 3–4 adjacent trees whose canopies are linked. They typically show limited movement, remaining in one spot for days and even weeks, as expected of a sit-and-wait predator. Adults feed on small birds and mammals, while juveniles take lizards and frogs. One field observation was of a snake at Fig. 2. A large female climbing vegetation.


Bako: Biodiversity Between Land and the Sea BORNEAN GREEN PIT-VIPER

a tree canopy lunging at flying bats! Vipers, in general, are known to swallow prey 56% of their body mass, relatively the largest prey swallowed by any snake, whereas the largest prey devoured by non-viper is only about half of this.

Vipers can complete a strike under a second. They can lunge from a resting position, erect the fangs, inject venom into their prey and return to their original position within the blink of an eye. The prominent fangs are particularly long, comprising hollow teeth that rotate through remarkable angles. While there are a number of records of bites by this species, no fatalities have been recorded. Nonetheless, the species needs to be treated with caution, as its venom contains haemotoxin that can damage red blood cells, in addition to a mixture of procoagulants that may have anticoagulants and/or haemorrhagic properties.

In Bako National Park, the Bornean Keeled Pit-viper is commonly encountered near the jetty, and provides tourists with a photographic opportunity.

Fig. 3. An adult about to strike.


The Editors

Jayasilan Mohd-Azlan (left) earned his doctoral degree from Charles Darwin University for his work on mangrove avifauna of Australia. He is currently the Director of the Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak.

Mohamad Kadim Suaidi (middle) is the Vice Chancellor of Universiti Malaysia Sarawak since 2013, and hails from Kampung Bako. He is passionate about community engagement and sustainable development in relation to biodiversity conservation.

The university’s visibility and recognition at the global stage is one of his main achievements.

Indraneil Das (right) received his doctoral degree from the University of Oxford, and was a Fulbright Fellow at the Museum of Comparative Zoology, Harvard University. Currently, he is Professor at the Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak.


This work takes the readers through a journey through several unique ecosystems within Bako National Park, highlighting many inhabitants on the way, from humble insects, such as mosquitoes and dragonfl ies, to the charismatic species, including the Proboscis Monkey and a rich assemblage of shorebirds that draw tourists by the thousands to the Park.

Bako is rich in biodiversity and accessible throughout the year via a short boat ride to the Park headquarters. From this vantage point, an eager tourist will be able to easily access the various habitats represented, including mixed dipterocarp forests, mangrove forests, cliff forest, beach forests, Kerangas, as well as mudfl ats, each with its unique biodiversity. The Park is also home to several endemic species, as well as species of conservation importance, upon which substantial ecotourism activities are based.

Bako National Park’s rich ecosystems are presented through images and text accounts in this volume, which is based on fi eld research, that reiterates their value for naturalists, tourists, as well as researchers. This book aims to enlighten stakeholders and present information on species biology and distribution to nature enthusiasts.

The chapter on geology and geomorphology sets the scene for the book.

The wildlife aspects cover species from an array of taxa that includes plants and invertebrates, to fi shes, herpetofauna, birds and mammals, rounded up with the ecotourism potential of the Park.

Research in Bako National Park was possible thanks to the generosity of various government agencies through research grants to Universiti Malaysia Sarawak. The project is aimed at assessing the area’s biotic diversity, examine anthropogenic elements, and fi nally, to develop an applicable environmental model for ecotourism.




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